Appetite 105 (2016) 591e595

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Appetite
journal homepage: www.elsevier.com/locate/appet

The evolutionary psychology of hunger

Laith Al-Shawaf a, b, *
a
Department of Psychology, Bilkent University, Ankara, Turkey
b
College of Life Sciences, Wissenschaftskolleg zu Berlin (Institute for Advanced Study), Germany

a r t i c l e i n f o a b s t r a c t

Article history: An evolutionary psychological perspective suggests that emotions can be understood as coordinating
Received 24 February 2016 mechanisms whose job is to regulate various psychological and physiological programs in the service of
Received in revised form solving an adaptive problem. This paper suggests that it may also be fruitful to approach hunger from this
14 June 2016
coordinating mechanism perspective. To this end, I put forward an evolutionary task analysis of hunger,
Accepted 15 June 2016
Available online 18 June 2016
generating novel a priori hypotheses about the coordinating effects of hunger on psychological processes
such as perception, attention, categorization, and memory. This approach appears empirically fruitful in
that it yields a bounty of testable new hypotheses.
Keywords:
Evolutionary psychology
© 2016 Elsevier Ltd. All rights reserved.
Hunger
Eating
Coordinating mechanisms
Superordinate mechanisms

1. Theoretical background opportunity (Al-Shawaf et al., 2015).

1.1. Emotions as coordinating mechanisms
An evolutionary psychological approach to the emotions sug-
gests that they are coordinating mechanisms e information-
processing programs that evolved to regulate the activity of other
programs in the service of solving a specific adaptive problem (Al-
Shawaf, Conroy-Beam, Asao, & Buss, 2015; Cosmides & Tooby,
2000; Tooby & Cosmides, 1990; 2008). On this view, the emotions
coordinate the activity of a wide range of psychological and phys-
iological mechanisms, ranging from perception, attention, and
memory to specialized inference mechanisms, physiology, and
behavior.
This approach suggests that each emotion evolved to coordinate
the operation of psychological and physiological mechanisms in
solving a particular adaptive problem. For example, fear co-
ordinates the activity of psychological and physiological programs
in the service of avoiding predators or escaping danger (Bracha,
2004; Marks & Nesse, 1994), disgust regulates mechanisms in or-
der to avoid infection (Curtis, Aunger, & Rabie, 2004; Tybur,
Lieberman, Kurzban, & DeScioli, 2012), and sexual arousal orches-
trates programs in preparation for an advantageous sexual
This coordinating mechanism approach was presented as a way
of conceptualizing emotions (Cosmides & Tooby, 2000; Tooby &
Cosmides, 1990; 2008), but it may also be usefully applied to
drives or motivational states. Hunger is usually regarded as a
motive or drive as opposed to an emotion (e.g., Kleinginna, Jr. &
Kleinginna, 1981; Lazarus, 1991; Oatley, Keltner, & Jenkins, 2006),
but it may be possible to use this approach to shed light on hunger
and to generate novel hypotheses about how it coordinates other
mechanisms in the body and mind.1 According to this view, hunger
may usefully be regarded as a mechanism that coordinates the
activity of psychological processes in the service of solving the
adaptive problem of acquiring food (Cosmides & Tooby, 2000).
In their presentation of the coordinating mechanism approach,
Tooby and Cosmides (1990, 2008; Cosmides & Tooby, 2000) sug-
gested a list of 14 programs regulated by the emotions. I present
these again here because they serve as a foundation for our hy-
pothesis generation method: (1) perceptual mechanisms, (2)
attention, (3) memory, (4) categorization, (5) motivational prior-
ities, (6) current goals, (7) information-gathering adaptations, (8)
specialized inference mechanisms, (9) communication & expres-
sion, (10) learning mechanisms, (11) reflexes, (12) energy level,

1
Indeed, the original authors of this idea suggest that, on their view, there may
* Department of Psychology, Bilkent University, Ankara, Turkey. be no principled, non-arbitrary distinction between drive states and other emotion
E-mail address: laith@bilkent.edu.tr. programs (Cosmides & Tooby, 2000).

http://dx.doi.org/10.1016/j.appet.2016.06.021
0195-6663/© 2016 Elsevier Ltd. All rights reserved.
592 L. Al-Shawaf / Appetite 105 (2016) 591e595
mood, and effort allocation, (13) physiology, and (14) behavior.
The theoretical foundations of this approach, including a dis-
cussion of a) the conceptual strengths of this view, b) why mech-
anisms need coordination in the first place, and c) how to
empirically test the validity of this approach, are all outlined in
longer conceptual papers (Al-Shawaf et al., 2015; Cosmides &
Tooby, 2000; Tooby & Cosmides, 1990; 2008). I therefore do not
repeat those points here. Instead, in this article I approach hunger
from this “coordinating mechanism” perspective, using a method
called evolutionary task analysis to generate an array of novel a
priori hypotheses about hunger.
1.2. Heuristic value and predictive power
The coordinating mechanism approach offers a systematic hy-
pothesis generation method based on evolutionary task analysis
(Marr, 1982). It consists of several key questions: (1) what adaptive
problem, if any, did this mechanism evolve to solve?, (2) which
subtasks must be solved in the solution of this adaptive problem?,
(3) which information-processing programs are capable of solving
these subtasks?, and (4) how should these programs be coordi-
nated to deliver a well-designed solution to this adaptive problem?
(Al-Shawaf et al., 2015).
This framework provides a systematic way to generate novel
empirical hypotheses for each candidate emotion or motivational
drive by asking which programs it is expected to coordinate and
how it is expected to do so. Of course, not every coordinating
mechanism is expected to regulate the full list of 14 programs listed
above.
In what follows, I apply this hypothesis-generation framework
to hunger. The purpose of this exercise is not to provide a
comprehensive analysis of hunger or generate an exhaustive list of
hypotheses, but rather to illustrate the heuristic value and predic-
tive power of this approach and to suggest that there may be utility
in conceptualizing hunger as a coordinating mechanism.
2. Novel hypotheses
2.1. Motivational priorities
Hunger, a universal mechanism that evolved to solve a critical
adaptive problem, is the first step in the food cycle e the sequence
of behaviors that culminates in eating (Rozin & Todd, 2015). In or-
der to solve the problem of food acquisition, hunger likely reorders
an individual’s motivational priorities, elevating the importance of
finding food and temporarily inhibiting other concerns such as
status enhancement, mating opportunities, and pathogen avoid-
ance. The intensity of this motivational reordering should depend
on the intensity of the hunger as well as the immediacy and
importance of other adaptive problems that the organism is facing.
For example, when simultaneously presented with hunger and a
threat or opportunity of great importance to fitness, hunger may
have a more muted effect on motivational reordering. In extreme
cases, intense hunger may be suppressed in order to solve a more
pressing adaptive problem, such as urgent predator avoidance or
the consummation of a valuable mating opportunity. We can refer
to the simultaneous presentation of conflicting adaptive problems
as an adaptive metaproblem.
Solving an adaptive metaproblem requires a tradeoff between
the mechanisms responsible for solving the competing adaptive
problems. In some cases, one adaptive problem may dominate, as
when hunger is suppressed during a near-death escape from a
predator. In other cases, a compromise solution is possible, and an
individual may simultaneously allocate a portion of her resources
to two different adaptive problems (see Crabbe, 2007; Prescott,
Bryson, & Seth, 2007; and Redgrave, Prescott, & Gurney, 1999, for
related discussions about natural action selection mechanisms).
Evolutionary psychological reasoning suggests that the solu-
tions to these adaptive metaproblems should differ systematically
by sex, context, and individual difference variables. For example,
given men’s stronger predilection for short-term mating and casual
sex (Buss, 2003; Lippa, 2009; Symons, 1979), a short-term mating
opportunity with an attractive member of the opposite sex should
suppress hunger much more powerfully among men than among
women. By contrast, given the robust sex difference in disgust
(women are more easily disgusted than men; Al-Shawaf & Lewis,
2013; Al-Shawaf, Lewis, & Buss, 2014; Curtis et al., 2004; Tybur
et al., 2012), the presentation of a pathogen threat should suppress
hunger more effectively among women than among men.
Ultimately, the outcome of tradeoffs between competing
mechanisms such as hunger and sexual arousal will depend on the
costs and benefits associated with allocating resources to one
adaptive problem over another. Ancestrally, the costs and benefits
of solving one adaptive problem over another would often have
differed systematically as a function of a) context, b) sex, and c)
individual difference variables such as immune competence, mate
value, and ability to evade predators.
These systematic differences in costs and benefits between
different contexts and individuals can be used to generate a priori
hypotheses about a) how hunger is expected to coordinate mech-
anisms in a context-specific and individual-specific manner, as well
as b) how hunger is expected to trade off with competing drives or
emotions when an organism is faced with distinct adaptive prob-
lems at once. More broadly, this coordinating mechanism approach
offers a way to generate context-specific hypotheses about how
hunger might regulate mechanisms in the body and mind.
2.2. Perception
Hunger may influence perceptual mechanisms in order to solve
the adaptive problem of finding food. Early theorists recognized
this possibility: “What we must study … are the variations
perception itself undergoes when one is hungry, in love, in pain, or
solving a problem” (Bruner & Goodman, 1947, p. 33).
These effects may operate through more than one perceptual
mode. For example, there is some evidence that hunger leads to
more positive evaluations of flavor, motivating further eating when
an individual has yet to reach satiation (e.g., Cabanac, 1971). We
might hypothesize that the gustatory reward associated with
eating should be directly dependent on hunger level, with greater
pleasure accompanying greater hunger and diminishing hedonic
returns accompanying increased satiety. Tying gustatory pleasure
to metabolic need in this way would be a good design for a system
that motivates eating.
There is already evidence to suggest that food odors e but not
other odors e are more pleasant when nutritionally depleted
(Duclaux, Feisthauer, & Cabanac, 1973). We also have evidence that
hunger increases the perceived brightness of foods (Gilchrist &
Nesberg, 1952), and improves perceptual encoding or accuracy of
food-relevant stimuli (Lazarus, 1998; Radel & Cle
ment-Guillotin,
2012), and that thirst may increase the perceptual or cognitive
accessibility of drinking-related stimuli (Aarts, Dijksterhuis, & De
Vries, 2001), as well as making it easier for people to perceive
transparency (a quality of water) in ambiguous stimuli (Changizi &
Hall, 2001). It would also be reasonable to expect that hunger and
thirst make food and drink look more visually appealing as well
(but see Hoefling et al., 2009).
Relatedly, hunger may enhance olfactory acuity, aiding the
process of detecting and locating food when nutritionally depleted.
This modulation of olfactory sensitivity by metabolic status and
 L. Al-Shawaf / Appetite 105 (2016) 591e595
feeding state has been demonstrated in nonhuman species such as
rodents, Drosophila, and C. elegans (Fadool, Tucker, & Pedarzani,
2011; Palouzier-Paulignan et al., 2012; Root, Ko, Jafari, & Wang,
2011). Testing this hypothesis in humans seems both feasible and
worthwhile.
An intriguing further possibility is that hunger may improve
detection of food-related sounds (e.g., sounds emitted by prey an-
imals, or the sound of group members eating nearby). These hy-
potheses about olfactory and auditory perception can be tested in
the lab by experimentally manipulating hunger and then present-
ing audio recordings and odors of different kinds, testing partici-
pants’ accuracy and sensitivity in detecting food-relevant and food-
irrelevant sounds and smells.
2.3. Attention and problem solving
Hunger may influence attention, too, narrowing an organism’s
attentional span and sharpening its focus to food and food-relevant
stimuli (Channon & Hayward, 1990; Mogg, Bradley, Hyare, & Lee,
1998; Stockburger, Schma €lzle, Flaisch, Bublatzky, & Schupp,
2009). To deliver a well-designed solution to this adaptive prob-
lem, hunger must down-regulate attention to other stimuli that
would normally capture an organism’s interest. Severe hunger may
even suppress attention to fitness-relevant stimuli such as patho-
gens, attractive mates, or valuable alliance-building opportunities
(but see Ainsworth & Maner, 2014).
It is also possible to generate very specific a priori predictions
about the effects of context and individual differences. For example,
hunger’s ability to shift attention away from pathogens and toward
food should be attenuated in people who are immunocompro-
mised. Hunger’s ability to suppress attention to other stimuli of
adaptive significance should also depend on the magnitude of the
fitness threat or opportunity posed by those other stimuli. For
example, hunger should be more effective at shifting a person’s
attention away from an unattractive mate or a moderately
dangerous predator compared to a highly attractive mate or an
extremely dangerous predator.
It may also be difficult for hungry individuals to sustain atten-
tion on food-irrelevant tasks. For example, hungry people may find
it hard to engage in future planning, problem solving, and extended
conversations. There should be a striking exception for food-
acquisition problems, which we might predict will elicit sustained
interest and attention from hungry individuals. I would therefore
hypothesize that hunger has opposite effects on different kinds of
problem solving: it leads to a decrement in people’s problem-
solving abilities in most domains, yet simultaneously enhances
their ability to solve food-acquisition problems.
We can test these predicted effects of hunger on attention and
problem solving using problem-solving tasks that vary in hunger-
relevance, dot-probe tasks, fMRI studies, and eye-tracking
methods (e.g., Mogg et al., 1998; Nijs, Muris, Euser, & Franken,
2010; Siep et al., 2009). As with many of the mechanisms dis-
cussed in this paper, the extent of hunger’s regulatory effects on
attention should depend on the intensity of the hunger.
2.4. Categorization and conceptual frameworks
Just as fear automatically imposes conceptual frameworks that
cause us to view our surroundings in terms of safety and danger
(Tooby & Cosmides, 1990), hunger may automatically impose
conceptual frameworks and categorization schemes relevant to the
adaptive problem of finding food. For example, at the most global
level, hunger may lead organisms to categorize items as either food
or not food, with different implications for approach behavior.
Within the category of food, items may be further categorized as
593
safe or spoiled. And nested in the category of safe foods, hunger may
further categorize items according to caloric density or nutritional
value. Higher priorities should be assigned to foods of greater value
and foods containing specific nutrients that an organism lacks (e.g.,
Rozin & Vollmecke, 1986; Rozin & Todd, 2015; Siep et al., 2009).
Evolutionary thinking suggests that this categorization may be
adaptively biased, such that intense hunger results in a more
lenient categorization of safe foods (e.g., see Cabanac, 1971). In the
extreme, we might expect an organism on the brink of starvation to
categorize as “safe” food sources that would otherwise be classified
as spoiled, as long as the on-average cost of such overinclusive
categorization is less than the cost of forgoing viable food sources
and risking death by starvation.
Extreme hunger should also suppress disgust to potentially
pathogenic foods (Hoefling et al., 2009). As individuals approach
starvation, we should see a muted disgust response to foods that
would otherwise trigger revulsion. If this response is an adaptation
against starvation, it should be highly specific to food, and not other
disgusting items. Evolutionary considerations therefore suggest
that hunger should have no effect on disgust toward open sores,
bodily effluvia, or other sources of pathogens and parasites. This
appears to be exactly the pattern of results we see (Hoefling et al.,
2009).
Importantly, severe food shortages were likely more common
among our hunter-gatherer ancestors than they are in today’s
industrialized nations, and hunger probably played a prominent
role in regulating psychology and behavior during human evolu-
tion. It is easy to underestimate the significance of this psycho-
physiological state in affluent modern-day societies, but hunger
remains a pervasive aspect of hundreds of millions of people’s lives
in developing nations and in modern hunter-gatherer societies
(Gat, 2000; Gurven, Hill, Kaplan, Hurtado, & Lyles, 2000; Sanchez &
Swaminathan, 2005; Serageldin, 2002; Speth & Spielmann, 1983),
and was likely a recurrent condition during the evolution of our
species.
In addition to affecting the way we categorize foods and path-
ogens, hunger may also affect the way we categorize other humans.
For example, hunger may lead us to categorize others as either
having or not having surplus food that they might be willing to
share. Similarly, hungry individuals may temporarily categorize
others in terms of whether they are likely to be useful in helping
with food acquisition (e.g., as a hunting or foraging partner).
The key point is that we may expect hunger to impose con-
ceptual frameworks and categorization schemes relevant to the
problem of securing food. We might therefore expect hunger to
cause people to adaptively categorize objects, other people, and
events in ways that facilitate the acquisition of food.
2.5. Memory
An intriguing possibility is that hunger might impact memory
mechanisms in the service of acquiring food. For example, hunger
may increase the accessibility of memories specifying locations in
which food was successfully obtained in the past. Indirect support
for this hypothesis comes from fMRI evidence that hunger prefer-
entially enhances recognition memory for food over nonfood
(Morris & Dolan, 2001), and from evidence that people in a field
setting show preferential recall for the locations of calorie-dense
food items over those of calorie-sparse ones (New, Krasnow,
Truxaw, & Gaulin, 2007; Krasnow et al., 2011). Additional evi-
dence that hunger might impact memory comes from research
showing superior word sequence recall for words directly following
food-related words, but only among hungry individuals (see Lerner,
Singer, & Triandis, 1958). During times of nutritional scarcity,
hunger may trigger especially strong encoding of information that
594 L. Al-Shawaf / Appetite 105 (2016) 591e595

can be used to resolve current or future hunger. Such a mechanism baseline activity in an orderly and predictable sequence, with more
might encode locations where food is available, useful methods of urgent and more impactful changes taking precedence. For
acquiring or preserving food, or individuals who offer food during example, the refocusing of attention on rivals or predators and the
one’s time of metabolic need. Intriguing indirect evidence for this reordering of motivational priorities may be more pressing than the
sort of hunger-mediated memory comes from studies showing that negation of hunger’s effects on memory. If this hypothesis is cor-
Drosophila flies “must be hungry to form and retrieve appetitive rect, we should expect to see broadened attentional focus and
memory” (Plaçais & Preat, 2013, p.440). renewed pathogen disgust before observing the reemergence of
We might also speculate that hunger could affect an individual’s memories of other duties and responsibilities. The broader point is
memory for group members. For example, hunger may temporarily that it is reasonable to expect the dissipation of hunger to sys-
increase the accessibility of memories indexing who typically tematically deactivate mechanisms in a predictable sequence ac-
shares food and who is stingy with it, who was recently observed to cording to the urgency or importance of the changes.
have surplus food, and who owes one an outstanding favor that At present, these suggestions are still speculative. But they
might be repaid in the currency of food. These hypotheses are highlight the fact that an evolutionary psychological approach
preliminary, but they represent an initial foray into the ways that points to a new conceptualization of hunger, novel hypotheses
hunger might regulate cognitive processes in the service of food about both hunger and eating, and new ideas about how eating
acquisition. And they are testable. may terminate the coordinating effects of hunger in a systematic
and predictable manner. Whatever its theoretical advantages, this
perspective is empirically fruitful in the sense that it provides a
2.6. Eating, the deactivation of hunger, and mechanism “offlining”
method for systematically generating a priori hypotheses about
hunger, as well as other drives and emotions.
Eating marks the solution of the adaptive problem that hunger
This approach also provides a logical way to generate hypoth-
was designed to solve. Eating should therefore be a critical stopping
eses about the “stopping points” of a drive or an emotion. When-
point, terminating the coordinating effects of hunger on other
ever a discrete event marks the solution to an adaptive problem
mechanisms (assuming that one eats enough to sate general hun-
that a motivational drive evolved to solve, that event can be ex-
ger plus any nutrient-specific hungers). This suggests the hypoth-
pected to cause the drive to dissipate. Consequently, the event in
esis that eating triggers a relatively abrupt deactivation of hunger’s
question should dissolve the coordinating effects of the drive,
coordinating effects on other mechanisms. This should manifest
resulting in a pronounced pre-event to post-event shift in a variety
itself in large pre-eating to post-eating shifts in people’s perception,
of mechanisms ranging from memory to attention to motivational
attention, categorization, memory, and motivational priorities.
priorities. We have hypothesized that in the case of sexual arousal,
For example, this hypothesis suggests that after sating them-
this “dissolving” event is orgasm (for males more strongly than
selves, people’s motivational priorities are once again reordered,
females; Al-Shawaf et al., 2015). In the case of hunger, this “dis-
attention is broadened beyond food-related stimuli, memories of
solving” event is eating. This leads to the testable hypothesis that
other duties and responsibilities resurface, categorization is no
eating until sated will cause the relatively rapid dissolution of
longer dominated by food, and previously suppressed adaptive
hunger’s effects on attention, perception, memory, and other psy-
problems such as mating and pathogen avoidance come to the fore
chological processes described above.
again. For example, the sight and smell of more food can become
disgusting after one is sated, a stark reversal in hunger’s disgust-
3. Conclusions
suppression effects. We might therefore expect that eating, a
discrete event that ushers in a suite of psychophysiological changes
An evolutionary psychological approach suggests that the
(Liu, Gao, & Fox, 2000), annuls or reverses the effects of hunger and
emotions are coordinating mechanisms whose evolved function is
marks the endpoint of the motivational state.2 This hypothesis
to orchestrate the activity of other mechanisms in the service of
suggests that we should see stark differences in people’s attention,
solving an adaptive problem (Al-Shawaf et al., 2015; Cosmides &
perception, categorization, and memory before and after eating. An
Tooby, 2000; Tooby & Cosmides, 2008). For example, disgust co-
intriguing hint of this possibility comes from a memory study:
ordinates mechanisms in the service of avoiding parasitic infection
“Most notably, in the 20 min interval between pre-satiation and
(Curtis et al., 2004; Tybur et al., 2012), fear regulates programs to
post-satiation scans, a >20% memory advantage for food stimuli
escape from danger (Bracha, 2004; Marks & Nesse, 1994; Marks,
relative to non-food was abolished in all 10 subjects” (Morris &
1987), and romantic love coordinates mechanisms in the service
Dolan, p. 5307).
of forming a long-term mating bond (Buss, 2006).
Of course, the effects of eating cannot be instantaneous. It takes
This approach might also be usefully applied to states such as
time for hunger to dissipate (e.g., Liu, Gao, Liu, & Fox, 2000; de
hunger, pain, and sexual arousal. These states are not traditionally
Graaf, Blom, Smeets, Stafleu, & Hendriks, 2004), so the return to
regarded as emotions, but may nonetheless be fruitfully regarded
baseline triggered by a meal may be speedy but not immediate. This
as coordinating mechanisms that have disparate effects on
return to baseline activity may be more pressing for certain
perception, attention, memory, conceptual categorization, and
mechanisms than for others. Stated differently, different sequences
other psychological processes.
of mechanism “offlining” may have different impacts on fitness.
Empirically, this approach provides a powerful hypothesis-
During the evolution of human hunger mechanisms, designs that
generation method. Researchers can use evolutionary task analysis
coordinated this “offlining” in a systematic and beneficial manner
to systematically generate an array of novel hypotheses for any
would have outcompeted designs that turned programs off
coordinating mechanism e including hunger e by asking (1) what
randomly or in inappropriate sequence.
adaptive problem, if any, the mechanism might have evolved to
This suggests the hypothesis that eating reverts mechanisms to
solve, (2) what subtasks constitute the adaptive problem, (3) which
psychological mechanisms are capable of solving these subtasks,
2
and (4) how these mechanisms ought to be coordinated to produce
This is admittedly an oversimplification. Eating is not a discrete event, but
rather a continuous process whose duration can vary. The “dissolving” effect of
a well-designed solution to the adaptive problem. This systematic
eating may therefore itself be continuous, and should depend on whether the or- approach can be applied to any candidate emotion or drive, pro-
ganism has eaten enough to sate itself. ducing a priori hypotheses about the effects of the mechanism on
 L. Al-Shawaf / Appetite 105 (2016) 591e595
psychological programs, the sequence of these effects, the context-
sensitivity of these effects, and even the coordinated dissolution of
these effects. Evolutionary task analysis thus offers both heuristic
value and predictive power, and may be used to great effect in the
study of emotions and motivational drives.More broadly, whatever
its theoretical merits, this “coordinating mechanism” approach
raises new questions, suggests new domains of inquiry, and ap-
pears to be empirically fruitful (Al-Shawaf et al., 2015; Cosmides &
Tooby, 2000; Tooby & Cosmides, 2008). The next step will be to test
some of the hypotheses in this paper and see how they stand up to
empirical investigation.
Acknowledgements
The author wishes to thank Peter M. Todd and an anonymous
reviewer for their helpful comments on an earlier draft of this pa-
per, as well as Dan Conroy-Beam for our early discussions about
adaptive metaproblems.
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